JUDITH GOODENOUGH, from the Department of Biology at the University of Massachusetts, has studied biological rhythms in creatures from planaria to finches to deer mice, and even in the green alga Chlamydomonas.

BETTY MCGUIRE is from the Department of Ecology and Evolutionary Biology at Cornell University. Her research focuses on parental behavior, reproduction, and ecology of small mammals, and she occasionally dabbles in work with larger domestic animals such as dogs and horses.

ELIZABETH JAKOB, from the Department of Psychology at the University of Massachusetts, studies the behavior of spiders, asking questions about their learning, perception, and interactions with conspecifics and with other species. She has carried out field projects in California, Mexico, Massachusetts, and Maine.

Chapter 1 Introduction 1

Four Questions About Animal Behavior 2

Animal Behavior as an Interdisciplinary Study 2

The Interplay of Questions: A Case Study 3

Hypothesis Testing 5

Part 1 Approaches to the Study of Animal Behavior

Chapter 2 History of the Study of Animal Behavior 11

The Beginnings 11

Intellectual Continuity in the Animal World 11

Darwin's Evolutionary Framework 12

Classical Ethology 13

The Approach: Evolutionary, Comparative, Descriptive, Field-Oriented 13

Classical Ethological Concepts 15

Comparative Psychology 17

The Approach: Physiological, Developmental, Quantitative, Laboratory-Oriented 17

Early Concepts of Comparative Psychology 17

The Roots of Physiological Psychology 20

Sociobiology and Behavioral Ecology 20

More Recent Trends 22

Field Studies 22

Cellular and Molecular Bases of Behavior 22

Behavioral Biology 23

Applied Animal Behavior 23

Chapter 3 Genetic Analysis of Behavior 27

Basics of Gene Action 28

Goals of Behavioral Genetics 31

Methods of Behavioral Genetics 31

Inbreeding 31

Artificial Selection 33

Inducing Mutations and Screening for Change in Behavior 35

Finding Natural Variants and Looking for Genetic Differences 36

Hybridization 37

The Foraging Gene as an Example of Behavioral Genetics in Action 38

Candidate Genes 38

Linking a Protein to a Trait 40

Locating all the Genes Associated with a Trait 41

Microarray Analysis 41

Important Principles of Behavioral Genetics 42

One Gene Usually Affects Several Traits 42

Genes Work in Interacting Networks 44

Behavioral Variation and Genes 45

Environmental Regulation of Gene Expression 47

Dominance Relationships in Cichlid Fish 47

Song Learning in Male Songbirds 48

The Importance of Genetic Background to Behavioral Genetics 50

Networks of Genes are Responsive to the Environment 51

Epigenetics and Behavioral Genetics 52

Complex Relationships Among Genes 54

A Broader Perspective 54

Chapter 4 Natural Selection and Behavior 57

Natural Selection 58

Common Misunderstandings About Natural Selection 60

Genetic Variation 61

Variation is Common 61

The Raw Material of Genetic Variation 61

Variation and the Response to Natural Selection 62

The Maintenance of Variation 63

Gene Flow and Genetic Drift 63

Correlated Traits 65

Changing Environmental Conditions 65

Frequency-Dependent Selection 67

Negative-Assortative Mating 68

Evolutionarily Stable Strategies: Fitness and the Behavior of Others 68

Testing Hypotheses About Natural Selection and Adaptation 70

The Experimental Approach 72

The Comparative Approach 73

Monitoring Selection in the Field 75

Modeling the Costs and Benefits of Traits 75

Chapter 5 Learning and Cognition 77

Definition of Learning 78

Types of Learning 78

Habituation 78

Classical Conditioning 80

Operant Conditioning 82

Latent Learning 84

Social Learning 85

Species Differences in Learning: Comparative Studies 87

The Ability to Learn as a Heritable Trait 87

Evolution and the Variation in Learning Across Species 88

Other Evidence of Cognitive Abilities in Animals 91

Tool Use 92

Detours 94

Understanding Numbers and Other Abstract Concepts 95

Self-Recognition and Perspective Taking 97

Chapter 6 Physiological Analysis-Nerve Cells and Behavior 99

Concepts from Cellular Neurobiology 100

Types of Neurons and Their Jobs 100

The Message of a Neuron 101

Ions, Membrane Permeability, and Behavior 104

Behavioral Change and Synaptic Transmission 104

The Structure of the Synapse 104

Integration 106

Specializations for Perception of Biologically Relevant Stimuli-Sensory Processing 107

Processing of Sensory Information for Sound Localization 108

Predators and Prey: The Neuroethology of Life-and- Death Struggles 109

Processing in the Central Nervous System 113

Brain Changes Underlying Behavioral Change 113

Social Behavior Network 120

Responding-Motor Systems 122

Neural Control in Motor Systems 122

Locust Flight 122

Chapter 7 Physiological Analysis of Behavior-The Endocrine System 127

The Endocrine System 129

Endocrine Glands and Hormones 129

Hormonal Versus Neural Communication 129

Types of Hormones and Their Modes of Action 130

How Hormones Influence Behavior 132

Effects on Sensation and Perception 132

Effects on Development and Activity of the Central Nervous System 133

Effects on Muscles 133

Methods of Studying Hormone-Behavior Relationships 135

Interventional Studies 135

Correlational Studies 137

Organizational and Activational Effects of Hormones 139

Defining the Dichotomy 139

Sex Differences in the Behavior of Norway Rats 139

Individual Differences in the Behavior of Male Tree Lizards 141

Questioning the Dichotomy 141

The Dynamic Relationship Between Hormones and Behavior 142

A Reciprocal Relationship 142

Hormonal Suppression of Behavior 142

Interactions Between Hormones, Behavior, and the Environment 142

Adjusting to the Harshness and Predictability of the Physical Environment 143

Adjusting to Onlookers in the Social Environment 145

A Detailed Look at the Hormonal Basis of Selected Behaviors 146

Helping at the Nest 146

Scent-Marking 147

Migrating 148

Chapter 8 The Development of Behavior 151

Influences on Behavioral Development 152

Development of the Nervous System 152

Development of Nonneural Structures 155

Hormonal Milieu 155

Physical Characteristics of the Environment 157

Experience Through Play 158

The Concept of Sensitive Periods 159

Changing Terminology-From Critical Periods to Sensitive Periods 159

Timing of Sensitive Periods 160

Multiple Sensitive Periods 161

Some Examples of Sensitive Periods in Behavioral Development 161

Pulling It All Together-The Development of Bird Song 171

Genetic, Hormonal, and Neural Control of Song 172

Role of Learning in Song Development 174

Sensitive Periods in Song Learning 176

Own-Species Bias in Song Learning 178

Social Factors and Song Development 178

A Diversity of Song Learning Strategies 179

Developmental Homeostasis 180

Rehabilitation of Chimpanzees After Long-Term Isolation 180

Part 2 Survival

Chapter 9 Biological Clocks 185

Defining Properties of Clock-Controlled Rhythms 186

Persistence in Constant Conditions 186

Entrainment by Environmental Cycles 187

Temperature Compensation 188

Rhythmic Behavior 189

Daily Rhythms 189

Lunar Day Rhythms 189

Semilunar Rhythms 189

Monthly Rhythms 190

Annual Rhythms 191

The Clock Versus the Hands of the Clock 192

Advantages of Clock-Controlled Behavior 192

Anticipation of Environmental Change 192

Synchronization of a Behavior with an Event That Cannot Be Sensed Directly 193

Continuous Measurement of Time 194

Adaptiveness of Biological Clocks 194

Organization of Circadian Systems 194

Multiple Clocks 195

Coordination of Circadian Timing 195

Human Implications of Circadian Rhythms 200

Jet Lag 200

Human Health 200

Chapter 10 Mechanisms of Orientation and Navigation 203

Levels of Navigational Ability 204

Piloting 204

Compass Orientation 204

True Navigation 206

Multiplicity of Orientation Cues 206

Visual Cues 206

Landmarks 206

Sun Compass 207

Star Compass 211

Polarized Light and Orientation 212

Magnetic Cues 214

Cues from the Earth's Magnetic Field 214

Directional Information from the Earth's Magnetic Field: A Magnetic Compass 217

Positional Information from the Earth's Magnetic Field: A Magnetic Map? 218

Magnetoreception 222

Chemical Cues 224

Olfaction and Salmon Homing 224

Olfaction and Pigeon Homing 226

Electrical Cues and Electrolocation 229

Chapter 11 The Ecology and Evolution of Spatial Distribution 233

Remaining at Home Versus Leaving 234

Costs and Benefits of Natal Philopatry 234

Costs and Benefits of Natal Dispersal 235

Sex Biases in Natal Dispersal 238

Natal Dispersal and Conservation Biology 240

Habitat Selection 241

Indicators of Habitat Quality 241

Search Tactics 243

Effects of Natal Experience 243

Habitat Selection and Conservation Biology 244

Migration 245

Costs of Migration 246

Benefits of Migration 248

Migration and Conservation Biology 251

Chapter 12 Foraging Behavior 253

Obtaining Food 253

Suspension Feeding 254

Omnivory 254

Herbivory 254

Carnivory 256

Adaptations for Detecting Prey 259

Optimal Foraging 264

Diet Selection: A Simple Model 265

Deciding When to Leave a Patch: The Marginal Value Theorem 267

Adding Complexity and Realism 268

The Utility of Models 274

Chapter 13 Antipredator Behavior 275

Camouflage 277

Coloration Matching the Visual Background 277

Disruptive Coloration 280

Countershading 282